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Ecology of Cichla (Cichlidae) in Two Blackwater Rivers of Southern Venezuela

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Ecology of Cichla (Cichlidae) in Two Blackwater Rivers of Southern Venezuela Copeia,1997(4), pp. 690-696 Ecology of Cichla (Cichlidae) in Two Blackwater Rivers of Southern Venezuela KIRK o. WINEMILLER, DONALD C. TAPHORN, AND MIELLO BARBARINO-DuQUE We investigated the ecology of peacock cichlids (Cichla spp.), diurnal piscovores and important gamefishes, in the Rio Cinaruco (Rio Orinoco drainage of the south- ern Venezuelan llanos) during the dry seasons of 1992-1993 and the Rio Pasimoni (Rio Negro-Amazonas drainage) during early 1993. In the Cinaruco, three Cichla species partitioned available habitats: C. intermedin near structure in primary river channel within or near swift current, C. orinocensisin shallow water along the shore- line of lagoons or slow channel reaches, and C. temensisin deeper waters along sandbanks and steeper rocky banks of lagoons and the river channel. During the dry season, C. orinocensisand C. temensisfed heavily on small characiform fishes, and C. intermedin consumed a variety of fishes, including loricariid catfishes. Only two peacock cichlids, C. orinocensisand C. temensis,were found in the Pasimoni, a more acidic and less seasonal ecosystem. These two species were larger in the Pas- imoni compared with Cinaruco conspecifics; however, conspecifics from the two locations conformed to similar mass-length relationships. In both river systems, C. temensiswere significantly larger than congeners, and C. temensisconsumed larger prey than congeners in the Cinaruco. Examination of gonads and sizes of intra- ovarian oocytes indicated that reproduction by all five populations probably is ini- tiated during the late low water period and may continue into the flooding period. Investigamos la ecologia de los pavones (Cichlidae, genero Cichla), picivores diur- nos importantes en la pesca deportiva, en el Rio Cinaruco (cuenca del Rio Orinoco en el sur del estado Apure, Venezuela) durante la sequia de los alios 1992-1993, y del Rio Pasimoni (cuenca Rio Amazonas, estado Amazonas) de Venezuela durante los primeros meses del 1993. En el Cinaruco, tres especies de Cichla reparten los habitat disponibles: C. intermedin se ubica cerca de cobertura (arboles sumergidas, etc.) en el cauce principal cerca 0 dentro de la corriente rapida; C orinocensisse encuentra mayormente en agua de poca profundidad cerca de la orilla de lagunas 0 en el cauce principal sin mucha corriente; y C. temensisocurre en agua mas pro- funda allado de bancos de arena y en lagunas y el cauce principal donde la orilla tiene una pendiente alta y es rocosa ("ripio 0 arrecife" de apurelio). Durante la sequia, C. orinocensisy C. temensisse alimentaron principalmente de pequelios peces caracidos, y C. intermedin consumi6 una variedad de peces, incluyendo corronchos loricoridos. Solamente dos pavones C. orinocensisy C. temensis,se encontraron en el rio Pasimoni, un ecosistema con menos fluctuaci6n estacional y mas acido. Estas dos especies alcanzaron tamanos mas grandes en el Pasimoni compararon con los pavones del rio Cinaruco de la misma especie, sin embargo, su curva tallo-peso era la misma. En ambas cuencas, los C. temensiseran significativamente mas grande que los otros pavones. En el rio Cincaruco, los C. temensisconsumieron presas mas gran- des que las de los otros dos pavones. Examinaci6n de la g6nades y los tamanos de los huevos indica que la reproducci6n en las cinco poblaciones de pavones estudia- dos probablemente se inicia durante la parte final de la sequia (aguas bajas) y con- tinua durante la etapa de lluvias e inundacion. P EACOCK cichlids (Cichla spp., Cichlidae) cies from the Amazon Basin require description are one of the most widespread and abun- (Kullander and Nijssen, 1989). Up to three co- dant predatory fishes in clearwater and black- existing Cichla species have been found in the water ecosystemsof the Orinoco and Amazon upper Rio Orinoco and upper Rio Negro basins Basins. Currently, systematists recognize five in southern Venezuela (Machado-Allison, 1971). Cichla species (known as pavon in Venezuela Peacock cichlids are diurnal piscivores that and'3s tucunare in Brazil), but additional spe- consume a variety of prey and potentially can f) 1997 by the American &JCiety of Ichthyologist, and HerpetoK'gists WINEMILLER ET AL.-ClCHIA ECOLOGY IN VENEZUELA 691 O"LaJ Fig. 2. Map showing the locations of the Rio Cin- aruco and Rio Pasimoni in southern Venezuela. Fig. 1. (A) The banded peacock cichlid, Cichla Ie- of these important Neotropical piscivores is merl.5is(81.5 cm SL), from the Rio Pasimoni; (B) royal based more on nonindigenous populations in peacock cichlid, Cichla intennedia (39 cm SL), from reservoirs than on fishes in their native fluvial the Rio Cinaruco; and (C) butterfly peacock cichlid, environments. We examined local population Cichla orinocensis(43 cm SL), from the Rio Pasimoni. structure, body mass-body length relationships, age at length, reproductive condition, patterns of habitat use, and diets of sympatric Cichla spe- influence prey populations. For example, the cies in two rivers of southern Venezuela. During introduction of exotic peacock cichlids into the dry season, Cichla ecology was investigated Lake Garon, Panama, was followed by massive in a river draining the southern savannas (low changes in community composition and the llanos) and a river that drains forest watersheds structure of the aquatic food web (Zaret and in southernmost Venezuela. Three species (C. Paine, 1973). A similar scenario of community intermedia, C. orinocensis,C. temensis,Fig. 1) are transition followed the introduction of C. ori- sympatric in the Rio Cinaruco (savanna). The nocensisand C. temensisinto Las Majaguas (AB-D, same three species coexist in rivers of forested Universidad Experimental de Los Llanos Occi- regions to the south (Machado-Allison, 1971), dentales thesis, unpubl., 1986; Winemiller, but C. intermediaapparently is absent from our 1989), Guri (Novoa et al., 1989; Gil et al., 1993), Rio Pasimoni study site. and other reservoirs in Venezuela. Zaret (1980) described the life history of a METHODS AND STUDY SITES nonindigenous population of C. ocellarisin Lake Garon, Panama. Characteristics of nonindigen- The Rio Cinaruco in southern Apure State ous C. ocellaris populations in Hawaiian and (R. Orinoco Basin) and the Rio Pasimoni in Puerto Rican reservoirs were summarized in southern Amazonas State of Venezuela (R. technical reports (W. Devick, Hawaii DLNR re- Casiquiare-R. Negro-R. Amazonas Basin; Fig. 2) port, unpubl., 1972; C. Lilyestrom et al., Puerto are low-gradient, blackwater rivers. The Cina- Rico DNER report, unpubl., 1994), and the life ruco originates in the flatlands of eastern Co- history of Cichla in fish farms was investigated lombia and flows eastward into Venezuela where in Brazil (Fontenele, 1950; Braga, 1952, 1953). it forms the southern boundary of Santos Lu- Very limited information on Cichla ecology in zardo National Park. The Cinaruco lies within a native rivers has been published (Lowe-Mc- region that experiences markedly seasonalrain- Connell, 1969; Goulding, 1980; Winemiller et fall and contains sandy, nutrient-poor soils, sa- al., 1995). Taphorn and Barbarino-Duque vanna vegetation, and acidic streams. In south- (1993) performed a mark/recapture study of ern Amazonas, the landscape is dominated by Cichla populations in rivers and lagoons of San- evergreen tropical forests. The Pasimoni origi- tos Luzardo National Park in Venezuela. nates on the northern slope of the Neblina 'Currently, our understanding of the ecology mountain range and flows northward to the Rio 692 COPEIA, 1997, NO.4 TABLE 1. WATER QUALITY PARAMETERSFOR THE Rio PASIMONI AND Rio CINARUCO (5-12 FEBRUARY1993). Rio Pasimoni Rio Cinaruco , Channel Lagoon Channel Lagoon pH 4.6 5.1 5.7 5.8 Conductivity(ltmhos/cm) 30 29 35 35 Total alkalinity (mg/l) <0.5 <0.5 <0.5 <0.5 Hardness(mg/l) <0.5 <0.5 6.0 6.0 Chloride (mg/l) 5.6 5.8 1.5 2.8 Color (ftu) 36 39 4 8 Total inorganic nitrogen (ltg/I) 29.3 30.0 7.0 11.2 Total inorganic phosphorus(ltg/I) 3.7 1.8 0.9 2.7 Temperature 26.0 26.0 30.0 31.0 Casiquiare and is surrounded by low, poorly growth checks on saggital otoliths were deter- drained terrain. Annual rainfall is greater, less mined for a few large individuals (C. temensis-- seasonal, and less predictable in this rain forest Pasimoni n = 4, C. temensi.r-Cinarucon = 3; region than in the llanos. We performed in ten- C. orinocensis--Cinaruco n = 2; C. intermedia-- sive surveys of the Cinaruco during 20-24 jan- Cinaruco n = 2). Recent research at the Cina- uary 1992, 7-12 February 1993, and 31 March ruco (D. jepsen, Texas A&M University, unpubl. to 2 April 1993; we surveyed the Pasimoni dur- thesis, 1995) indicates that growth checks are ing 31 january to 5 February 1993. annuli that form during the late dry season The following environmental parameters (March to April) when fish prepare for spawn- were estimated using meters (YS1, Inc.): con- ing. Most Cichla specimens were handled in the ductivity, salinity, temperature, and dissolved ox- field, but a few were preserved in formalin and ygen. Standard titration methods were used to returned to the Universidad Experimental de measure total alkalinity, chloride, total hard- Los Llanos Occidentales or Texas A&M for ness, pH, water color, and total dissolved inor- stomach contents analysis. ganic nitrogen and phosphorus; and visual es- timates were used to record substratum com- position, submerged woody structure, channel RESULTS width, water depth, and relative current velocity. Fishes were sampled in the river channel and Water samples from the Pasimoni and Cina- lagoons with seines, gillnets, castnets, and hook ruco revealed classic blackwater features (Sioli, and line. Angling with lures produced almost all 1984) of low pH, conductivity, solute, and nu- of our Cichla specimens (SL > 20 cm), and al- trient concentrations (Table 1). Water chemis- though no juvenile Cichla were captured, a few try varied little between lentic lagoon and lotic juveniles were observed in shallow backwaters channel habitats within each river.
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